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Accepted by M.R. de Carvalho: 27 Feb. 2015; published: 25 Mar. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3937 (2): 347
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http://dx.doi.org/10.11646/zootaxa.3937.2.6
http://zoobank.org/urn:lsid:zoobank.org:pub:E654E1ED-708D-4537-A7EC-8FFF8EC2DCF4
Scyliorhinus ugoi, a new species of catshark from Brazil (Chondrichthyes:
Carcharhiniformes: Scyliorhinidae)
KARLA D. A. SOARES
1,4
, OTTO F. B. GADIG
2
& ULISSES L. GOMES
3
¹Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Rua do Matão, Trav. 14, no. 101, São Paulo, SP, CEP
05508-900, Brazil. E-mail:
karlad.soares@yahoo.com.br
²Laboratório de Pesquisa de Elasmobrânquios, Universidade Estadual Paulista ‘Júlio Mesquita Filho’, Campus Experimental do
Litoral Paulista, Departamento de Biologia Marinha, Praça Infante Dom Henrique, s/n, Parque Bitaru, São Vicente, SP, CEP 11330-
900, Brazil. E-mail: gadig@clp.unesp.br
³Laboratório de Taxonomia de Elasmobrânquios, Universidade do Estado do Rio de Janeiro, Instituto de Biologia,
Departamento de
Zoologia, Rua São Francisco Xavier, 524, Maracanã, Rio de Janeiro, RJ, CEP 20550-900. E-mail: ulisseslg@uerj.br
4
Corresponding au thor
Abstract
A new species of catshark (Carcharhiniformes, Scyliorhinidae), Scyliorhinus ugoi sp. nov., is described from off North-
eastern and Southeastern Brazil. The new species is closest to the Scyliorhinus haeckelii/besnardi group and S. hesperius
but differs in background coloration, head width, sexual maturity, and in cranial and body proportions.
Key words: Scyliorhinidae, Scyliorhinus ugoi sp. nov., dark freckled catshark, southwestern Atlantic Ocean, Brazil
Introduction
The genus Scyliorhinus Blainville 1816, consists of 15 species whose geographical distribution covers all seas,
from cold to tropical waters, except the Antarctic and Southeastern Indian Ocean. Species of this genus live in close
association with the substrate, from coastal regions to the edge of the continental shelf, reaching depths of
about 2,000 m (Bigelow & Schroeder, 1948; Springer, 1966, 1979; Compagno et al., 2005).
According to Springer (1979) and Compagno (1984), Scyliorhinus presents the following characters: color
pattern with dark dorsal saddles, clear and/or dark blotches, supraorbital crest present, small snout, small lower
labial furrow and upper labial furrow absent, and second dorsal fin smaller than first. There is a small flap that
slightly overlaps the lower lip at each corner of the mouth, the outer edge of which may extend anteriorl y,
from the corner of the mouth for a short distance, giving the impression that an upper labial furrow is present.
Scyliorhinus is most diverse in the western Atlantic and at least three species are listed as occurring in the
Brazilian coast, as follows: S. haeckelii Miranda Ribeiro, 1907, distributed from Venezuela to Argentina
(Compagno, 1984; Figueiroa, 2011), S. besnardi Springer & Sadowsky, 1970, found in southeastern Brazil to
Argentina (Springer & Sadowsky, 1970, Springer, 1966, 1979; Figueiroa, 2011), and S. hesperius Springer, 1966,
recorded from Honduras in the Caribbean Sea to the Brazilian central coast (Gadig & Gomes, 2003). But because
of their great similarity and lack of exclusive characters, S. haeckelii and S. besnardi are treated here as the S.
haeckelii/besnardi group (Gadig, 2001; Gomes et al., 2010; Soares, 2014). However, Soares (2014) has found that
S. hesperius of Gadig & Gomes (2003) and Scyliorhinus sp. of Gomes et al. (2010) represent a new species
occurring from northeastern Brazil to Rio de Janeiro state; this species is herein described.
Material and methods
Abbreviations and methodology for body measurements followed Compagno (2001) with addition of the
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measurement ‘vent-caudal length’. Measurements are expressed as percentages of total length (%TL). Cranium
terminology and cranial measurements followed Compagno (1988a). Clasper terminology was based on
Jungersen (1899). Counts of v ertebrae and fin radials were t a k en from radiographs and dissected specimens.
Tooth row counts and nomenclature for dentition followed Compagno (1988a). Dermal denticles were taken from
the left side of the body above the pectoral and below first dorsal fins. Dermal denticles were photographed
using a scanning electron microscope (SEM) at the Universidade do Estado do Rio de Janeiro (UERJ). The
distribution of the new species is shown in a map which was generated based on the software Google Earth.The
holotype and paratypes (Fig. 1) were deposited in the Museu Nacional, Rio de Janeiro (MNRJ), Museu de
Zoologia da Universidade de São Paulo (MZUSP) and ichthyological collection of the Universidade do Estado do
Rio de Janeiro (UERJ).
Scyliorhinidae Gill, 1862
Scyliorhinus Blainville, 1816
Scyliorhinus ugoi sp. nov.
(Figs. 1–9)
Proposed common name: dark freckled catshark (English), cação-gato negrinho (Portuguese).
Scyliohinus sp.: Gadig, 2001, pp. 149–150, coast of Espírito Santo state; Gomes et al. 2010, pp. 85–86, fig. 110, coast of Rio de
Janeiro state.
Scyliorhinus hesperius: Gadig & Gomes, 2003, p. 22, coasts of Espírito Santo and Rio de Janeiro states.
Scyliorhinus sp. 1: Soares, 2014, pp. 80–84, figs. 17, 68.
Holotype. MNRJ 42619, adult female, 496 mm TL (southern Bahia, Central Brazilian coast).
Paratypes. MZUSP 110448, adult male, 465 mm TL (Alagoas, Northeastern Brazil, 9° S 34° 50’W);
MZUSP 110449, adult male, 445 mm TL (Rio Grande do Norte, Northeastern Brazil, 6° 14’S 34° 51’W); UERJ
1725, adult male, 530 mm TL (Central Brazilian coast, between southern Bahia and northern Rio de Janeiro;
cranium, jaws and claspers); UERJ 2179, immature male, 415 mm TL (southern Bahia, Central Brazilian coast).
Additional material examined. 5 specimens (see Appendix).
Diagnosis. A southwestern Atlantic Ocean Scyliorhinus distinguished by a considerably broad head (vs.
slender in the S. haeckelii/besnardi group and S. hesperius), and color pattern with brown saddles on a light
brown background, predorsal saddles with anterior and posterior sharp median projections (vs. no projections
in the S. haeckelii/besnardi group or S. hesperius); and light and dark, spiracle-sized or slightly smaller spots,
lacking large white spots (vs. large white spots in S. hesperius). The following combination of characters also
distinguishes these species: snout rounded and moderately short, preoral length 5% TL (vs. 4.5% TL in the S.
haeckelii/besnardi group); preorbital length 6.4% TL and head length 20% TL (vs. 5% TL and 18% TL in S.
hesperius); mouth width corresponding to 9% TL (vs. 8% TL in the S. haeckelii/besnardi group and 7.2% TL in S.
hesperius); pectoral-pelvic space 1.5 times pelvic-anal space (vs. equal in the S. haeckelii/besnardi group), and
interdorsal space 2.3 times dorsal-caudal space (vs. 1.6 times in S. haeckelii/besnardi group); anal-fin base 8% TL
(vs. 6.7% TL in S. hesperius and 8.3% TL in the S. haeckelii/besnardi group); cranium with flat basal plate, without
keels; cranial width larger in females (42.7% NL) (vs. 37.4% NL in the S. haeckelii/besnardi group); a terminal
dermal cover present in hindmost portion of clasper, covering terminal cartilages (lacking in S. hesperius);
medium-sized, adult male at 445 mm TL and adult female 500 mm TL (vs. 353 mm and 410 mm, respectively, for
the S. haeckelii/besnardi group).
Description. Morphometric measurements are given in Table 1 and cranial measurements in Table 2. Modes of
meristic counts are given as well their range in parentheses (if different).
Head (Fig. 2a–c) broad and depressed, greatest width 0.63 times of head length. Snout thick, somewhat
flattened and relatively short, preoral length corresponding to 4.3–5.6% TL and 0.5 times the mouth width.
Ampullae pores not greatly enlarged on snout. Preoral length 1.1–1.4 times in preorbital length (vs. 1.4–1.7 in the
Scyliorhinus haeckelii/besnardi group) (Tab. 1).
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DESCRIPTION OF SCYLIORHINUS UGOI
TABLE 1. Measurements in millimeters and proportions as percentage of total length for Scyliorhinus ugoi sp.
nov. Mean (x), standard deviation (SD) and range of proportions (%).
Measurements Scyliorhinus ugoi sp. nov. Scyliorhinus haeckelii/besnardi
x SD% x SD %
Total length (TL) 516 ±68.1 100 354.4 ±78.7 100
Precaudal length 393.7± 55.7 74.7–76.8 275.6 ±63.5 76.5–77.9
Eye-spiracle length 6.1 ± 1.3 0.9–1.3 3.3 ±1.2 0.6–1.3
Preoral length 23.8 ±4.7 4.3–5.6 16.9 ±3.2 4.3–4.7
Preorbital length 32.6 ±3.8 6.4 23.2 ±4.5 6.2–8.0
Prespiracular length 53.1 ±5.9 10.4–10.7 37.6 ±7.7 10.6–10.7
Prebranchial length 76.6 ±11.4 14.5–15.2 54.2±11.9 14.8–15.4
Head length 103.2 ±14.3 19.5–20.3 71 ±16.1 18.7–19.2
Prepectoral length 94.0 ±12.1 17.9 66 ±15.3 16.1–18.2
Prepelvic length 217.5±33.8 41.0–43.4 148 ±33.5 38.2–43.7
Snout-vent length 232.2±34.9 43.6–45.4 152 ±33.9 40.9–42.3
Vent-caudal length 262.5 ±43.4 53.7–55.3 189.6 ±40.9 51.3–58.3
Preanal length 314.1 ±50.3 59.1–64.4 213.2 ±51.4 55–62.7
Pre-first dorsal length 257.5 ±38.2 48.4–51.2 177 ±41.3 44.9–51.3
Interdorsal space 57.2 ±9.5 10.9–11.0 40.4 ±13.5 7.0–10.7
Dorsal-caudal space 22.3 ±4.7 4.2– 5.1 18.2 ±3.9 5.8–6.7
Pectoral-pelvic space 96.3 ±16.4 17.6–20.1 63.5 ±15.9 10–20.3
Pelvic-anal space 59.7 ±13.3 10.5–12.8 44.1 ±12.3 12.7–14.6
Anal-caudal space 39.2 ±5.6 7.4–7.6 28.5 ±6.5 8.7–8.9
Eye length 19.8 ±2.8 3.8–4.0 13.6 ±3.4 3.3–4.3
Eye height 3.8 ±0.3 0.9–1.4 4.7 ±1.1 1.3–1.5
Internarial space 10.1 ±3.6 1.4–2.4 9 ±3.4 2.0–4.0
Interorbital space 31.1 ±3.1 6.1–6.6 24.3 ±4.8 6.4–7.3
Spiracle length 4.3 ±1.1 0.4–0.9 2.6 ±1.1 0.6–1.3
Mouth length 23.5 ±5.0 3.7–5. 1 14.9 ±4.8 4.7–5.6
Mouth width 46.6 ±7.4 9.0–9.7 27.8 ±6.4 8.0
Lower labial furrow length 10.6 ±1.9 1.9–2.1 6.6 ±1.8 1.3–2.0
First gill slit height 15.2 ±2.7 2.8–3.1 9 ±2.3 2.0–2.8
Fifth gill slit height 8.6 ±2.3 0.9–1.8 4.7 ±1.3 1.3–1.9
Pectoral length 68.5 ±7.4 12.8–13.6 41 ±11.4 10.7–13.8
Pectoral anterior margin 74.6 ±9.0 14.5–15.0 47.5 ±11.9 12.7–15.6
Pectoral base 34.1 ±10.3 3.1–7.2 24.3 ±7.1 6.7–9.7
Pectoral posterior margin 45.6 ±6.3 8.7–9.0 30 ±8.8 5.3–8.9
Pectoral inner margin 32.2 ±2.3 5.9–7.1 21.9 ±5.8 5.3–7.6
Pelvic length 54.3 ±10.1 9.0–10.5 37.58 ±11.4 8.7–12.5
Pelvic inner margin 18.0 ±3.1 3.1–3.8 16.3 ±6.7 4–6.8
Pelvic base 38.0 ±11.0 6.4–8.9 24.4 ±6.6 4.7–8.0
Pelvic anterior margin 36.0 ±5.8 6.6–7.4 23 ±6.1 5.3–7.0
Pelvic posterior margin 33.6 ±4.8 6.4–6.6 22.7 ±7.2 3.2–7.4
Clasper outer length 20.3 ±4.5 3.8–4.1 16.8 ±8.1 2.0–6.2
......continued on the next page
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Eyes (Fig. 3) large and spindle-shaped; eye length 5.2 times in head length and 2.5–4.2 times eye height.
Eyes dorsolateral on head, with lower edges well medial to horizontal head rim in dorsal view; subocular ridges
strong. Nictitating lower eyelids of rudimentary type (sensu Compagno, 1970), with shallow subocular pouches
and secondary lower eyelids free from upper eyelids. Prespiracular length corresponding to 10.5% TL (Tab. 1).
Nostrils (Fig. 4) with broad incurrent apertures, without nasoral grooves or nasal barbels, small oval excurrent
apertures and short posterior nasal flaps. Anterior nasal flaps large, covering posterior nasal flaps and
excurrent apertures somewhat anterior to mouth. Internarial space 2.4–4.6 times in interorbital space.Mouth (Fig.
4) broadly arched, moderately wide and short, mouth width 2.2 times in head length; mouth length 0.5 times
in mouth width. Teeth prominently exposed in ventral view. Lower labial furrows short, 0.2 times in mouth
width; upper labial furrows absent. Tongue moderate-sized flat and rounded, occupying most of mouth floor.
Dorsal labial cartilages 1.5 times the ventral one; anterior tip of dorsal cartilages reaching orbital processes of
palatoquadrate.
TABLE 1. (Continued)
Measurements Scyliorhinus ugoi sp. nov. Scyliorhinus haeckelii/besnardi
x SD% x SD %
Clasper inner length 44.5 ±14.8 8.1–9.0 35.7 ±12.6 6.0–10.4
Clasper base width 3.5 ±2.1 0.4–0.8 4 ±1.4 0.6–1.1
Anal base 39.8 ±8.0 5.7–7.7 30.2 ±6.3 8.0–8.7
Anal anterior margin 44.1±6.3 8.3–8.7 28.1 ±6.3 8.0–8.7
Anal posterior margin 26.8 ±3.1 5.1–5.4 17.6 ±4.9 4.7–6.0
Anal inner margin 13.5 ±2.3 2.6–2.9 9 ±3.7 2.0–4.9
Anal height 22.3 ±3.8 4.2–4.6 13 ±3.6 3.3–5.1
First dorsal anterior margin 51.1 ±6.3 9.7–10.0 34.9 ±8.5 8.7–10.6
First dorsal base 36.1 ±5.7 6.6–7.1 24.1 ±6.4 6.7–8.0
First dorsal height 30.2 ±4.9 5.7 –5.9 19.1 ±5.7 2.7–6.0
First dorsal inner margin 15.7 ±2.7 3.1–3.4 10.2 ±2.8 2.0–3.6
Second dorsal anterior margin 36.6 ±7.9 5.0–7.6 26.5 ±6.1 5.3–7.6
Second dorsal base 30.1 ±4.8 5.5–6.1 19.5 ±4.9 5.3–6.6
Second dorsal height 21.1 ±4.6 3.8–4.7 12.6 ±3.4 3.3–4.7
Second dorsal inner margin 13.7 ±2.5 2.6–2.9 9 ±2.4 1.3–3.0
Dorsal caudal margin 122.5 ±13.4 23.4–25.3 78.7 ±16 23.2–23.5
Postventral caudal margin 50.7 ±4.0 9.4–10.7 35.4 ±7.5 8.7–10.0
Subterminal caudal margin 48.5 ±9.0 8.8–10.7 30.9 ±6.8 8.0–8.5
Terminal caudal margin 25.8 ±4.0 5.2–5.4 18.4 ±5.1 0.6–7.0
Terminal caudal lobe 29.0 ±4.0 5.4–5.6 18.6 ±5.3 4.0–7.2
Head height 32.6 ±7.4 5.4–6.6 22.7 ±5.8 4.7–7.0
Trunk height 53.8 ±11.4 9.5–12.2 42.2 ±8.9 10.7–11.6
Abdomen height 37.3 ±7.0 7.1–8.2 28.5 ±8 6.0–8.9
Caudal height 28.8 ±8.3 5.0–7.6 20.9 ±6 5.3–7.4
Caudal peduncle height 15.8 ±2.5 3.1–3.3 12.6 ±3.8 3.3–4.9
Head width 65.0 ±8.6 13. 0–13.3 41.4 ±13.5 3.3–11.7
Trunk width 63.8 ±13.7 10.9–14.5 43.7 ±10.3 11.4–12.7
Abdomen width 42.3 ±9.2 6.6–8.7 26.7 ±7.2 6.7–8.9
Caudal width 22.0 ±5.0 3.8–4.9 16 ±4.2 4.0–4.9
Caudal peduncle width 12.0 ±2.2 1.9–2.3 10.8 ±3.6 2.7–4.1
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DESCRIPTION OF SCYLIORHINUS UGOI
FIGURE 1. Scyliorhinus ugoi sp. nov., holotype, MNRJ 42619, female, 500 mm TL. A) Dorsal view. B) Lateral view;
paratype, UERJ 2179, male, 419 mm TL. C) Dorsal view. D) Lateral view. Scale bar: 5 cm.
First two gill openings about equally long; first gill opening twice as large as fifth (Fig. 2c). All gill openings
slightly concave and not elevated on dorsolateral surface of head, gill filaments not visible externally.
Sexual heterodonty weak; adult males presenting teeth longer with undeveloped lateral cusps on central
portion of lower jaw. Upper teeth slightly higher-crowned than lower teeth, with longer, stronger transverse
ridges, otherwise very similar. Medial teeth (M) and anteroposterior teeth (AN) poorly differentiated; symphysial
teeth (S) with higher-crown and smaller than anteroposteriors, with erect or semi-erect cusps and one weak
cusplet on either side of the cusp. Anteroposteriors in both jaws larger than medials and symphysials, with semi
oblique cusps, usually one strong high cusplet on either side, longitudinal ridges confined to the basal ledges,
and low, flat roots. Gradient monognathic heterodonty well-developed in anteroposterior teeth; anteroposteriors
smaller distally, with thicker and more oblique cusps, and lower cusplets.
Teeth in 52/49 (47–56/45–53) rows, 3/2 (2–3/2–3) functional series. Teeth not highly differentiated in upper
and lower jaws or along jaws. Tooth formula:
Left AN16–19 M6–8 S1 S1 M6–8 AN16–19 Right
AN15–17 M6–9 M7–10 AN16–17
Lateral trunk denticles (Fig. 5) with flat, elongated teardrop-shaped crowns 1.4–2.1 times as long as wide,
anterior part covered with ectodermal pits (sensu Muñoz- Chápuli, 1985). Crown with a strong medial ridge
that extends entire length of crown on the long medial cusp. Lateral cusps not well developed, 0.3 times in medial
cusp; lateral ridges anterior to them very short or absent. Males with dermal denticles longer than females.
Density of dermal denticles on posterior regions lower than on anterior regions.
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TABLE 2. Measurements in millimeters and proportions as percentage of nasobasal length for Scyliorhinus ugoi sp. nov.
Female (F), male (M), and range of proportions (%NL).
Body (Fig. 1) stout and depressed on head region, tapering considerably to caudal fin. Prepectoral length
corresponding to 17.9% of total length (TL) and prepelvic length 41–43.4% TL. Snout–vent length 0.8 times in
vent-caudal length. Pectoral-pelvic space 1.5–1.6 times pelvic–anal space (vs. 0.8–1.4 in the Scyliorhinus
haeckelii/besnardi group). Interdorsal space 2.1–2.5 times dorsal-caudal space (Tab. 1).
Pectoral fins (Figs. 1 and 2) large and rounded-triangular, not falcate, with narrowly rounded apexes and broad
bases. Origin of pectorals under interspace between third and fourth gill openings. Pectoral base 0.3– 0.7 times
mouth width (vs. 0.8 times in the Scyliorhinus haeckelii/besnardi group). Pectoral anterior margin 2–4.8 times its
base (vs. 1.7–2.3 in the Scyliorhinus haeckelii/besnardi group) and 1.6 times the posterior margin (Tab. 1).
Pectoral fin skeleton aplesodic (Fig. 6) with radials mostly divided into three segments, longest distal
radial segment (DRA) 1.8 times length of its proximal radial segment (PRA). Pectoral skeleton tribasal,
propterygium (PRO) with a single radial, mesopterygium (MES) with 3 radials, and metapterygium (MET) with
8 ( 7–9) radials on basal segment and 1 ( 1–2) on metapterygial axis (MTS); total radial count 13 (12–15).
Propterygium small, short and rectangular or trapezoidal. Mesopterygium short, diamond-shaped and slightly
elongated and distally expanded in the axes of its radials. A fused plate formed by the proximal segment of the
second, third and fourth mesopterygial radials. Metapterygium basal segment triangular, elongated slightly
diagonal to the axes of its radials; metapterygial axis quadrangular, trisegmental, and with length about 0.4
times of metapterygial basal segment. This segment presents a projection directed anteriorly in its inner margin.
Pelvic fins (Fig. 2) broadly triangular; pelvic anterior margins 0.5 times of pectoral anterior margins. Pelvic
anterior margin 0.9–1.1 times posterior margin (Tab.1). Total radial count 13 (12–15).
Claspers (Fig. 7) moderately long and slender, cylindrical and blunt-tipped, extending free rear tips of pelvic
fins, by a distance about 0.2 times pelvic inner margin. Most of clasper except dorsomedial and posteromedial
surface of glans (including rhipidion) covered by large denticles with anteriorly directed cusps. An envelope
(EN) delimits anteriorly an exorhipidion (ERH) not differentiated from lateral edge of glans, specialized clasper
hooks absent. Rhipidion (RH) well developed medially covered by a poorly developed cover rhipidion (CRH).
Measurements Scyliorhinus ugoi sp. nov.
F % NL M % NL
Nasobasal length (NL)54.3 100 51.5 100
Rostral length 14.6 20.8–29.3 12 22.4–24
Width across lateral rostral cartilages 7.6 13.8–14.5 6.5 12.2–13
Width across nasal capsules 43.6 77.5– 85.4 40 76–79.6
Nasal capsule width 21.6 37.9–43.7 21 40.8
Nasal capsule length 20.6 37.5–39.6 19 35.1–38.7
Nasal aperture width 13.5 25–25.8 14 26.5–27.7
Distance between nasal apertures 6.6 10.4–13.8 6.5 12.2–13
Distance between rostral base and anterior edge of anterior
fontanelle
21 39.5 19.5 36.7–38.8
Anterior fontanelle width 13.3 24.1–25 12.5 22.4–26
Basal plate width 22.6 39.6–45.8 19.5 37–38.7
Orbit length 24.3 44.8–47.9 24 46.3–47
Preorbital process length 3.3 6.2–7.4 3.5 6.1–7.4
Postorbital process length 4.5 8.3–8.6 4.5 8.1–9.2
Nasal capsules length 11 18.7–20.7 10.5 20.4
Suborbital shelf width 36 65.5–73 32.5 62.8–63.2
Otic capsules width 28 54.1–57.4 28.5 55.1–55.5
Width across preorbital processes 31.6 56.9–62.5 29.5 57.1–57.5
Width across postorbital processes 39.5 72.4–77 36 69.4–70.3
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DESCRIPTION OF SCYLIORHINUS UGOI
Pseudosiphon poorly developed and visualized only internally; pseudopera absent. Apopyle (AP) and hypopyle
(HP) connected by a long clasper groove, with its dorsal margins fused over the clasper groove. Clasper skeleton
(Fig. 7b) relatively simple. The axial cartilage connected proximally by a single basal segment and a dorsal beta
cartilage to the pelvic basipterygium. Clasper shaft slender and formed from the axial cartilage and tightly
rolled dorsal and ventral marginal (RD and RV) cartilages. A poorly developed acessory dorsal marginal
cartilage (RD2) present between the terminal portion of the dorsal marginal cartilage and anterior portion of the
dorsal terminal (TD); this cartilage supports the rhipidion. The skeleton of the clasper glans comprises a large,
curved, wedge-shaped dorsal terminal and a similarly shaped ventral terminal (TV), articulating with and
separated by a terminal extension of the axial cartilage, the narrow end-style (G); ventral terminal longer than
dorsal terminal cartilage. There is no accessory terminal cartilage or dorsal terminal 2 cartilage, but a slender
ventral terminal cartilage (TV2) is positioned on the ventral terminal cartilage. A terminal dermal cover (TDC –
new term) present in the hindmost portion of the clasper, covering the terminal cartilages and presenting no
denticles.
FIGURE 2. Scyliorhinus ugoi sp. nov. A) Dorsal view of the head of a female, MNRJ 42619, female, 500 mm TL. B)
Dorsal view of the head of a male, UERJ 2179, 419 mm TL. C) Lateral view of head, UERJ 2179 male, 419 mm TL. D)
First
dorsal fin. E) Second dorsal fin. F) Anal fin. G) Caudal fin. H) Ventral view of trunk, MNRJ 42619, female, 500 mm
TL. Scale bars: 2 cm.
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FIGURE 3. Eye of Scyliorhinus ugoi sp. nov., MNRJ 42619, female, 500 mm TL. Scale bar: 2 cm.
FIGURE 4. Nostrils and mouth of Scyliorhinus ugoi sp. nov., MNRJ 42619, female, 500 mm TL. Scale bar: 2 cm.
First dorsal fin (Figs. 1, 2) triangular, apically narrow and not falcate, with nearly straight anterior margin,
rounded apex and angular free rear tip. First dorsal origin above to insertions of pelvic fins. First dorsal base
1.6 times in interdorsal space, 3.4 times in dorsal caudal margin. First dorsal anterior margin 1.4 times its base;
first dorsal height 0.8 times first dorsal base. Total radial count 13 (12–14).
Second dorsal fin (Figs. 1, 2) low, triangular, apically narrow and not falcate; second dorsal fin height 0.7
times first dorsal fin height, base 0.8 times first dorsal fin base. Second dorsal fin with nearly straight
anterior margin, bluntly rounded apex, narrowly rounded free rear tip, and straight inner margin. Second
dorsal fin origin slightly behind anal midbase. Second dorsal fin base 0.7 times in dorsocaudal space, second
dorsal fin height 1.4 times in second dorsal fin base. Total radial count 10 (9–12).
Anal fin low (Figs. 1, 2), apically narrow, not falcate and somewhat larger than second dorsal fin, anal fin
height 1.1 times second dorsal fin height and base 1.3 times second dorsal fin base. Anal fin anterior margin
nearly straight, apex narrowly rounded, free rear tip acutely pointed, and inner margin straight. Anal fin base
without preanal ridges, anal fin origin and anal fin base length behind pelvic fins insertions. Anal fin base 0.5
times interdorsal space (vs. 0.6–0.8 in the Scyliorhinus haeckelii/besnardi group) and 1.3–1.5 times dorsocaudal
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space. Anal anterior margin 1.5–1.7 times posterior one, anal fin height 1.8 times in anal fin base. Total radial
count 20 (18–22).
Caudal fin (Figs. 1, 2) narrow-lobed and asymmetrical, with large and hardly developed terminal lobe.
Caudal fin short, dorsal margin 3.2 times in precaudal length; subterminal margin 0.9–1 time in terminal margin.
Dorsal caudal fin margin slightly convex, without lateral undulations. No crests of denticles on the caudal fin
margins.
FIGURE 5. Lateral trunk denticles of Scyliorhinus ugoi sp. nov. A–C) UERJ 1726, female, 602 mm TL. D–F) UERJ 1725,
adult male, 534 mm TL. A, D) Region above pectoral fin. B, E) Region below fi r s t dorsal fin. C, F) Region below
second dorsal fin.
FIGURE 6. Pectoral fin skeleton of Scyliorhinus ugoi sp. nov., UERJ 1722, female, 600 mm TL. Scale bar: 2 cm. DRA,
distal segment; MES, mesopterygium; MET, metapterygium; MTS, metapterygial axis; PRA, proximal segment; PRO,
propterygium.
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FIGURE 7. Clasper of Scyliorhinus ugoi sp. nov., UERJ 1725, male, 530 mm TL. A) Clasper gland. B) Skeleton. Scale bar =
5 mm. AP, apopyle; CRH, cover rhipidion; G, end-style; EN, envelope; ERH, exorhipidion; HP, hypopyle; RD, marginal dorsal
cartilage; RD2, accessory marginal dorsal cartilage; RH, rhipidion; RV, marginal ventral cartilage; TD, terminal dorsal
cartilage; TDC, terminal dermal cover; TV, terminal ventral cartilage; TV2, terminal ventral cartilage 2.
Total vertebral counts 127 (119–135), monospondylous precaudal centra (MP) 39 (38–39). Transition
between monospondylous and diplospondylous (DP) centra behind pelvic fin bases and over clasper shafts.
Last MP centra before MP–DP transition smaller than the anterior and larger than the first DP, not forming a
'stutter- zone' of alternating long and short centra.
Intestinal valve of conicospiral type, with 6 (6–8) turns.
Cranium (Fig. 8) broad and flattened, corresponding to 10% TL (Tab. 2). Rostral cartilages short and
slender; rostrum length 20.8–29.3% of nasobasal length (NL). Medial rostral cartilage (MR) originates from
anteroventral tip of internasal septum and extends anterodorsally. Distance between lateral rostral cartilages
(LR) proportionately larger on females (13.8–14.5% NL) than males (12.2–13% NL). Nasal capsules (NC)
large, transversely oval, its width larger than length. Width across capsules 76– 85.4% NL. Nasal apertures
(NA) broadly circular and laterally positioned on capsules; two thirds of its area covered by nasal fontanelle
(NF). Males with nasal apertures proportionately larger (26.5–27.7% NL) than females (25–25.8% NL).
Internasal septum (IS) high and compressed, not hypercalcified in adults. Distance between rostral base and
anterior edge of anterior fontanelle (AF) proportionately larger in females (39.5% NL) than males
(36.7– 38.8% NL). Cranial roof (CRF) broadly arched between supraorbital crests (SC) in frontal view, with a
definite saddle-like medial depression in lateral view. Anterior fontanelle broad and subquadrate on males and
heart-shaped on females, length 1.3 times width, width 4 times in nasobasal length. Basal plate flat, without keels;
width larger in females (39.6–45.8% NL) (vs. 33.3–40.8% NL in the Scyliorhinus haeckelii/besnardi group).
Orbits oval-subquadrate in shape; its length 2.2 times in nasobasal length. Supraorbital crests low, strong and
deeply arched. Postorbital processes (PTP) angular, situated on anterior ends of optic capsules; width across
postorbital processes 1.2 times preorbital processes (PEP). Optic capsules short, length 4.7 times in nasobasal
length; its width 2.5–2.7 times optic capsules length. Hyomandibular facets (HF) large, extending across the
entire lateral face of each optic capsule, with slightly exserted ends.
Color pattern with brown saddles on a light brown background, predorsal saddles with anterior and posterior
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sharp median projections; light and dark spots spiracle-sized or slightly smaller, no large white spots. Light brown
on ventral region; some specimens with dark points on body and fins. Remarkably darker than the Scyliorhinus
haeckelii/besnardi group and S. hesperius.
FIGURE 8. Cranium of Scyliorhinus ugoi sp. nov., UERJ 1725, male, 530 mm TL. A) Dorsal view. B) Ventral view; UERJ
1726, female, 597 mm TL. C) Dorsal view. D) Ventral view. Scale bar = 1 cm. AF, anterior fontanelle; ASC, anterior
semicircular canal; CRF, cranial roof; FOE, external foramen of preorbital canal; FPE, external profundus foramen; HF,
hyomandibular facet; ICF, internal carotid foramen; IS, internasal septum; LR, lateral rostral cartilage; MR, medial rostral
cartilage; NA, nasal aperture; NC, nasal capsule; NF, nasal fontanelle; OC, optic capsule; PEP, preorbital process; PRF,
parietal fossa; PSC, posterior semicircular canal; PT, pterotic process; PTP, postorbital process; SC, supraorbital crest;
SF, stapedial foramen; SG, supraorbital groove; SS, suborbital shelf.
Distribution and biological data. This species (Fig. 9) is recorded from the coast of the states of Rio Grande
do Norte, Pernambuco, Alagoas, Bahia (northeastern Brazil), Espírito Santo, and northern Rio de Janeiro
(southeastern Brazil). Accurate occurrence records are known only for a few specimens, but this species has been
collected for several years between Rio Grande do Norte and Rio de Janeiro states. Most specimens recorded are
demersal, about 400–500 m deep, associated with coral formations, where it possibly lays its egg-cases. Stomach
contents include small invertebrates (cephalopods), small bony fishes, and one hagfish (Class Myxini) egg-case
was found in the stomach of an adult specimen. Maximum total length about 630 mm. Size of the first maturity not
well defined, but 450 mm TL males and 470 mm TL females are adults. Males ranging in size from 419–534 mm
and females from 431–630 mm. Reproduction is by oviparity, producing at least two amber colored egg capsules,
one for each uterus, which are smooth, lacking longitudinal grooves and measuring about 152 mm long and 58 mm
wide (Gadig, 2001; Gomes et al, 2010).
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FIGURE 9. Distribution of Scyliorhinus ugoi sp. nov. Yellow diamonds represent records of type series, and white star
locality of the holotype.
Etymology. The specific name ugoi is dedicated to Ugo de Luna Gomes, son of Ulisses L. Gomes.
Comparisons with other species of Scyliorhinus. Seven species of Scyliorhinus are readily distinguished
from Scyliorhinus ugoi by color pattern. Scyliorhinus retifer (Garman, 1881) has a unique reticulated pattern of
dark lines; S. boa Goode & Bean, 1896, S. canicula Linnaeus, 1758, S. cervigoni Maurin & Bonnet, 1970, S.
garmani (Fowler, 1934), and S. stellaris Linnaeus, 1758 have conspicuous, small to medium-sized, rounded dark
spots and indistinct saddle blotches (Springer, 1979; Compagno, 1984; Compagno et al., 2005). In the Scyliorhinus
haeckelii/besnardi group, when contrasting saddles are present, specimens lack pre-dorsal saddles with anterior
and posterior sharp median projections (Soares, 2014). Scyliorhinus meadi Springer, 1966 has a color pattern of
dark saddles on a light background, without light or dark spots (Springer, 1979; Compagno, 1984).
Scyliorhinus ugoi, S comoroensis Compagno, 1988, S. capensis (Müller & Henle, 1838), S. torrei Howell-
Rivero, 1936, S. hesperius Springer, 1966, S. tokubee Shirai, Hagiwara & Nakaya, 1992, and S. torazame (Tanaka,
1908) are similar in having a color pattern of light spots on a darker background, combined with dark saddles, but
only Scyliorhinus ugoi presents small dark spots and pre-dorsal saddles with anterior and posterior sharp
median projections (Springer, 1979; Compagno, 1988; Shirai et al., 1992; Compagno et al., 2005; Soares, 2014).
Scyliorhinus canicula has relatively larger, broader anterior nasal flaps that reach the mouth, covering the
shallow nasoral grooves that are lacking in S. ugoi and in all other species of the genus (Springer, 1979;
Compagno, 1984; Compagno et al., 2005; Soares, 2014).
Scyliorhinus haeckelii/besnardi from the Southwestern Atlantic Ocean differs from S. ugoi in its smaller size,
with adult males known at about 350–370 mm, while in S. ugoi males at about 420 mm are still immature. Its
snout is more slender, usually more pointed and longer than that of S. ugoi, with preoral length 1.4–1.7 times in
preorbital length vs. 1.1–1 . 4 times in S. ugoi (Soares, 2014). The wider and stout head of S. ugoi is easily
observable in all specimens and, together with its characteristic color pattern, permits a precise distinction.
The abdomen of S. ugoi is relatively longer than in S. haeckelii/besnardi, with pectoral–pelvic space 1.5–1.6
times pelvic–anal space vs 0.8–1.4. Interdorsal space 2.1–2.5 times dorsal–caudal space vs. 1.2–2 in the
Scyliorhinus haeckelii/besnardi group (Soares, 2014).
Monospondylous vertebral counts of S. ugoi (38–39) differ from counts for S. meadi (43–49), S. torrei
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(30–34), S. comoroensis (45) and S. capensis (48), but do not differ appreciably from counts for S. retifer, S.
boa, S. hesperius and S. haeckelii/besnardi (Springer & Sadowsky, 1970; Springer, 1979; Compagno, 1988b;
Soares, 2014).
The occurrence of S. hesperius in the uppermost slope (274–457 m in depth) of the Atlantic from Central
America to Brazil (Gadig & Gomes, 2003) is an error of identification and refers, in fact, to S. ugoi (Soares,
2014). Beyond the differences in color pattern, S. ugoi has a larger head, with preorbital length 6.4% TL and
head length 19.5–20.3% TL vs. 4.9–5.3% TL and 18.9% TL in S. hesperius, respectively. The mouth is also
larger in S. ugoi, its width 9.0–9.7% TL vs. 7.0–7.5 in S. hesperius, as is its anal base length at 7.7–8.2% TL vs.
5.7–7.7% TL in S. hesperius (Springer, 1979; Soares, 2014).
It is apparent from the comparisons presented above that S. ugoi is not particularly close to its geographically
nearest congener, S. haeckelii/besnardi of the southwestern Atlantic Ocean, and is differentiated from S. hesperius
of Central America by color pattern and body proportions.
Acknowledgements
The authors wish to acknowledge José Lima Figueiredo, Aléssio Datovo (MZUSP) and Lynne Parenti (USNM)
for the loan of specimens used in this work. We also acknowledge Herculano Alvarenga (MHNT) for the
radiographs of some specimens and Alan Moraes (Laboratory of Eletronical Microscopy, UERJ) for help with the
SEM. Marcelo Carvalho (IBUSP) is thanked for extensive review of the manuscript. The first and second authors
are supported by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (Capes) and Conselho
Nacional de Desenvolvimento Científico e Tecnológico (CNPq), respectively.
References
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Compagno, L.J.V. (1970) Systematics of the genus Hemitriakis (Selachii: Carcharhinidae), and related genera. Proceedings of
California Academic Sciences, Series 4, 38, 63–98.
Compagno, L.J.V. (1984) FAO species catalogue. Vol. 4. Sharks of the world: An annotated and illustrated catalogue of
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Compagno, L.J.V. (1988a) Sharks of the Order Carcharhiniformes. Princeton University Press, 467 pp.
Compagno, L.J.V. (1988b) Scyliorhinus comoroensis sp. n., a new catshark from the Comoro Islands, western Indian Ocean
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Compagno, L.J.V. (2001) Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Vol.
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Compagno, L.J.V., Dando, M. & Fowler, S. (2005) Sharks of the world. Princeton University Press, Princeton, 368 pp.
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P.Y. & Cosseau, M.B. (Eds.), Contribuciones sobre biologia, pesca y comercialización de tiburones en la Argentina.
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Gadig, O.B.F. (2001) Tubarões da costa brasileira. Unpubl. PhD thesis, Universidade Estadual Paulista ‘Júlio Mesquita Filho’,
Rio Claro, São Paulo, 343 pp.
Gadig, O.B.F. & Gomes, U.L. (2003) Classe Chondrichthy es. In: Menezes, N.A., Buckup, P.A., Figueiredo, J.L. & Moura,
R.L. (Eds.), Catálogo das espécies de peixes marinhos do Brasil. Museu de Zoologia da Universidade de São Paulo,
São Paulo, pp. 21–22.
Gomes, U.L., Signori, C.N., Gadig, O.B.F. & Santos, H.R.S. (2010) Guia para Identificação de Tubarões e Raias do Rio de
Janeiro. Technical Books, Rio de Janeiro, 234 pp.
Jungersen, H.F.E. (1899) On the apendices genitales in the greenland shark Somniosus microcephalus (Bl. Schn.) and
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Shirai, S., Hagiwara, S. & Nakaya, K. (1992) Scyliorhinus tokubee sp. nov. from Izu Peninsula, Southern Japan (Scyliorhinidae,
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Soares, K.D.A. (2014) Estudo taxonômico das espécies de Scyliorhinus do grupo haeckelii/besnardi (Carcharhiniformes,
Scyliorhinidae) das regiões Sudeste e Sul do Brasil (Oceano Atlântico Sul Ocidental). Unpubl. Master's thesis,
Universidade do Estado do Rio de Janeiro, Rio de Janeiro, 208 pp.
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Springer, S. (1966) A review of Western Atlantic catsharks, Scyliorhinidae, with descriptions of a new genus and five new
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Biological Society of Washington, 83 (7), 83–98.
APPENDIX. Comparative material.
Scyliorhinus ugoi: 5 specimens. UERJ 1426, female, 513 mm TL (off Bahia, Central Brazilian coast); UERJ 1722, adult
female, 600 mm TL (off Salvador, Bahia, Central Brazilian coast); UERJ 1723, female, 427 mm TL (Central Brazilian coast,
between Pernambuco and northern Rio de Janeiro); UERJ 1726, adult female, 597 mm TL (Central Brazilian coast, between
Pernambuco and northern Rio de Janeiro); UNESP-CLP 0093, adult female, 580 mm TL (Bahia, Northeastern Brazil).
Scyliorhinus boa: 9 specimens. USNM 186195.1, male, 350 mm TL (6°29’N 52°30’W, French Guiana); USNM 186195.2,
male, 315 mm TL (6°29’N 52°30’W, French Guiana); USNM 186195.3, female, 285 mm TL (6°29’N 52°30’W, French
Guiana); USNM 204378, female, 435 mm TL (11º15’N 68°13’W, Caribbean Venezuela); USNM 221565.1, female, 425
mm TL (11º50’N 73°5’W, Caribbean Colombia); USNM 221565.2, female, 350 mm TL (11º50’N 73°5’W, Caribbean
Colombia); USNM 221565.3, male, 376 mm TL (11º50’N 73°5’W, Caribbean Colombia); USNM 221566, male, 210
mm TL (11°58’N 69°30’W, Caribbean Venezuela); USNM 221567, male, 205 mm TL (11°31’N 64°11’W, Caribbean
Venezuela).
Scyliorhinus haeckelii/besnardi: 92 specimens. MNRJ 494, male, 317 mm (Ilha Rasa, Rio de Janeiro, Southeastern Brazil);
MZUSP 9963, female, 329 mm TL (35°18'S 52° 32'W, Uruguay; MZUSP 9965, male, 361 mm TL (35°18'S 52° 32'W,
Uruguay); MZUSP 9966, male, 169 mm TL (35°18'S 52° 32'W, Uruguay); MZUSP 37282, female, 280 mm TL (23°05’S 41°
59'W, São Paulo, Southeastern Brazil); MZUSP 37283, male, 356 mm TL (23°05’S 41° 59'W, São Paulo, Southeastern
Brazil); MZUSP 37284, male, 391 mm TL (23°10’S 43º05’W, São Paulo, Southeastern Brazil); MZUSP 37284, female, 355
mm TL (23°10’S 43º05’W, São Paulo, Southeastern Brazil); UERJ 71.1, female, 394 mm TL (Southeastern Brazil); UERJ
71.2, male, 364 mm TL (Southeastern Brazil); UERJ 1489, male, 491 mm TL (Southeastern Brazil); UERJ 1574, female,
371 mm TL (off Paraná, Southern Brazil); UERJ 1496.1, female, 361 mm TL (Itajaí, Santa Catarina, southern Brazil); UERJ
1496.2, female, 367 mm TL (Itajaí, Santa Catarina, southern Brazil); UERJ 1573, female, 297 mm TL (off Paraná, southern
Brazil); UERJ 1689, male, 566 mm TL (Southeastern Brazil); UERJ 1690, female, 467 mm TL (Southeastern Brazil); UERJ
1691, male, 522 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 1692, male, 389 mm TL (Southeastern Brazil); UERJ
1693, male, 469 mm TL (Cabo Frio, Rio de Janeiro, Southeastern Brazil); UERJ 1695, female, 494 mm TL (Southeastern
Brazil); UERJ 1696, female, 451 mm TL (Southeastern Brazil); UERJ 1697, male, 491 mm TL (Southeastern Brazil);
UERJ 1698, male, 454 mm TL (Southeastern Brazil); UERJ 1699, female, 400 mm TL (Cabo Frio, Rio de Janeiro,
Southeastern Brazil); UERJ 1701, female, 405 mm TL (Cabo Frio, Rio de Janeiro, Southeastern Brazil); UERJ 1703, male,
451 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 1704, male, 425 mm TL (Southeastern Brazil); UERJ 1705, female,
450 mm TL (Cabo F rio, Rio de Janeiro, Southeastern Brazil); UERJ 1706, female, 422 mm TL (Cabo Frio, Rio de
Janeiro, Southeastern Brazil); UERJ 2202, male, 444 mm TL (Southeastern Brazil); UERJ 2208, male, 216 mm TL
(Southeastern Brazil); UERJ 2232.1, female, 341 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.2, female, 304
mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.3, female, 348 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ
2232.4, male, 383 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.5, male, 321 mm TL (Rio de Janeiro,
Southeastern Brazil); UERJ 2232.6, male, 341 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.7, male, 415 mm
TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.8, male, 337 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ
2232.9, male, 293 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.10, male, 318 mm TL (Southeastern Brazil);
UERJ 2232.11, female, 334 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.12, female, 326 mm TL (Rio de
Janeiro, Southeastern Brazil); UERJ 2232.13, male, 325 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.14,
female, 334 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.15, female, 305 mm TL (Rio de Janeiro,
Southeastern Brazil); UERJ 2232.16, male, 314 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.17, male, 296
mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.18, male, 274 mm TL (Rio de Janeiro, Southeastern Brazil);
UERJ 2232.19, male, 368 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.20, female, 323 mm TL (Rio de
Janeiro, Southeastern Brazil); UERJ 2232.21, male, 353 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.23, male,
416 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.24, female, 241 mm TL (Rio de Janeiro, Southeastern
Brazil); UERJ 2232.25, female, 326 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.26, female, 305 mm TL (Rio
de Janeiro, Southeastern Brazil); UERJ 2232.27, female, 277 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.28,
female, 308 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.33, male, 315 mm TL (Rio de Janeiro, Southeastern
Brazil); UERJ 2232.39, female, 299 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.7, female, 348 mm TL (Rio
de Janeiro, Southeastern Brazil); UERJ 2233.13, female, 366 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.14,
female, 250 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.29, female, 370 mm TL (Rio de Janeiro, Southeastern
Brazil); UERJ 2232.30, male, 344 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.31, female, 269 mm TL (Rio
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de Janeiro, Southeastern Brazil); UERJ 2232.32, male, 262 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.34,
female, 259 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.35, male, 339 mm TL (Rio de Janeiro, Southeastern
Brazil); UERJ 2232.36, male, 282 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.37, male, 382 mm TL (Rio
de Janeiro, Southeastern Brazil); UERJ 2232.38, female, 334 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ
2232.40, male, 341 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.41, female, 281 mm TL (Rio de Janeiro,
Southeastern Brazil); UERJ 2232.42, female, 255 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.43, male, 281
mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.45, female, 300 mm TL (Rio de Janeiro, Southeastern Brazil);
UERJ 2232.47, female, 255 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.49, female, 306 mm TL (Rio de
Janeiro, Southeastern Brazil); UERJ 2232.50, female, 318 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.51,
female, 287 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2232.52, female, 335 mm TL (Rio de Janeiro,
Southeastern Brazil); UERJ 2233.1, male, 353 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.2, male, 438 mm
TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.3, female, 409 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ
2233.4, female, 401 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.5, female, 357 mm TL (Rio de Janeiro,
Southeastern Brazil); UERJ 2233.6, male, 449 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.8, female, 443 mm
TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.9, female, 413 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ
2233.10, female, 432 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.11, male, 447 mm TL (Rio de Janeiro,
Southeastern Brazil); UERJ 2233.12, female, 461 mm TL (Rio de Janeiro, Southeastern Brazil); UERJ 2233.15, female, 413
mm TL (Rio de Janeiro, Southeastern Brazil).
Scyliorhinus hesperius: 1 specimen. USNM 187729, female, 265 mm TL (16°52’N, 81°30’W).
Scyliorhinus meadi: 1 specimen. USNM 187730, female, 459 mm TL (17°40’N, 77°55’W).
ABBREVIATIONS
AF, anterior fontanelle; AN, anteroposterior tooth; AP, apopyle; ASC, anterior semicircular canal; CRF, cranial
roof; CRH, cover rhipidion; DP, diplospondylous centra; DRA, distal segment; G, end-style; EN, envelope; ERH,
exorhipidion; FOE, external foramen of preorbital canal; FPE, external profundus foramen; HF, hyomandibular
facet; HP, hypopyle; IBUSP, Instituto de Biociências da Universidade de São Paulo; ICF, internal carotid foramen;
IS, internasal septum; LR, lateral rostral cartilage; M, medial tooth; MÊS, mesopterygium; MET, metapterygium;
MHNT, Museu de História Natural de Taubaté; MP, monospondylous centra; MR, medial rostral cartilage; MTS,
metapterygial axis; NA, nasal aperture; NC, nasal capsule; NF, nasal fontanelle; NL, nasobasal length; PEP,
preorbital process; PRA, proximal segment; PRF, parietal fossa; PRO, propterygium; PSC, posterior semicircular
canal; PT, pterotic process; PTP, postorbital process; RD, marginal dorsal cartilage; RD2, accessory marginal dorsal
cartilage; RH, rhipidion; RV, marginal ventral cartilage; S, symphysial tooth; SC, supraorbital crest; SF, stapedial
foramen; SG, supraorbital groove; SS, suborbital shelf; TD, terminal dorsal cartilage; TDC, terminal dermal cover;
TL, total length; TV, terminal ventral cartilage; TV2, terminal ventral cartilage 2; USNM, Smithsonian National
Museum of Natural History.
